Impact of native vegetation cover near crops on the occurrence and molecular diversity of fire ants

Solenopsis invicta and Solenopsis saevissima are the most common fire ants in Brazil. Environmental disturbances favour the colony dispersal of both species, particularly those with an anthropogenic origin. However, the frequencies of the species can vary according to the amount of anthropogenic disturbances. Crops near native vegetation fragments is a common landscape in the Brazilian Atlantic Forest. Here, we analyse if there is influence of native vegetation cover on the occurrence of these fire ants and their molecular diversity in crops. Our hypothesis is that these species are antagonistic in terms of occurrence and molecular diversity when compared in the same habitat. We selected crops near fragments with around 50% of native vegetation cover, a percentage informative enough to detect biological responses from insects belonging to a same order. Nests were collected and the species were identified using external morphology, and mtDNA COI sequences. We reject our hypothesis, which suggest that the species colonize the vegetation patches around crops in a similar way. Solenopsis invicta is not limited by vegetation cover, and has a high haplotype diversity when compared to S. saevissima, particularly when the vegetation cover is greater than 50%. Additionally, the vegetation cover seems to not have influence in the total number of haplotypes, but the species have haplotypes that are exclusive to each landscape. Our findings suggest that S. invicta is expanding to native vegetation areas that it has not occupied before, as S. saevissima was dominant there.     

The Guianese population of the fire ant Solenopsis saevissima is unicolonial

In this study, conducted in French Guiana, a part of the native range of the fire ant Solenopsis saevissima, we compared the cuticular hydrocarbon profiles of media workers with previous results based on intraspecific aggressiveness tests. We noted a strong congruence between the two studies permitting us to delimit 2 supercolonies extending over large distances (up to 54 km), a phenomenon known as unicoloniality. Solenopsis geminata workers, taken as an out‐group for cluster analyses, have a very different cuticular hydrocarbon profile. Because S. saevissima has been reported outside its native range, our conclusion is that this species has the potential to become invasive because unicoloniality (i.e., the main attribute for ants to become invasive) was shown at least for the Guianese population.     

SYSTEMATICS AND POPULATION GENETICS OF FIRE ANTS (SOLENOPSIS SAEVISSIMA COMPLEX) FROM ARGENTINA

Specimens of seven fire ant species collected from their native ranges in Argentina were studied by protein electrophoresis and morphological analysis. Concordance between the genetic and morphological character sets is strong (96% agreement on identifications), suggesting that recognition of reproductively isolated populations and partitioning of intra- and interspecific variation can in most cases be achieved using appropriate characters of either type in this taxonomically difficult group. Genetic differentiation between native (Argentina) and introduced (USA) conspecific populations of two species, Solenopsis invicta and S. richteri, is rather typical of the differentiation existing between conspecific populations found within either country. Furthermore, there appears to have been little reduction of variability (heterozygosity) at enzyme loci following colonization by either species of the United States, although some rare alleles have been lost in the introduced populations. Hybridization is rare between S. invicta and S. richteri where their native ranges overlap in central Argentina, in contrast to the extensive hybridization of these species in the United States, suggesting that prezygotic barriers to gene flow have been compromised in introduced populations. Phylogenetic analysis of the seven species indicates that S. invicta and S. richteri are relatively distantly related within the S. saevissima complex. Given that hybrids between these species in the United States suffer little apparent loss of fitness, genomic incompatibilities generally may be insufficient to create effective postzygotic barriers to interspecific gene flow in this group of ants.     

Traits allowing some ant species to nest syntopically with the fire ant Solenopsis saevissima in its native range

Supercolonies of the red fire ant Solenopsis saevissima (Smith) develop in disturbed environments and likely alter the ant community in the native range of the species. For example, in French Guiana only 8 ant species were repeatedly noted as nesting in close vicinity to its mounds. Here, we verified if a shared set of biological, ecological, and behavioral traits might explain how these 8 species are able to nest in the presence of S. saevissima. We did not find this to be the case. We did find, however, that all of them are able to live in disturbed habitats. It is likely that over the course of evolution each of these species acquired the capacity to live syntopically with S. saevissima through its own set of traits, where colony size (4 species develop large colonies), cuticular compounds which do not trigger aggressiveness (6 species) and submissive behaviors (4 species) complement each other.     

COMPARATIVE BIOCHEMICAL GENETICS OF THREE FIRE ANT SPECIES IN NORTH AMERICA,WITH SPECIAL REFERENCE TO THE TWO SOCIAL FORMS OF SOLENOPSIS INVICTA (HYMENOPTERA: FORMICIDAE)

An electrophoretic study of the genetics of three fire ant species in North America was undertaken with the primary objective of further clarifying the genetic relationship between two social forms of Solenopsis invicta. Such social forms are common in many groups of ants and may, in some cases, represent significant intermediate stages in the speciation process. The monogyne and polygyne forms of S. invicta, while differing in a number of important biological traits, are genetically indistinguishable, in contrast to the substantial genetic differentiation observed between S. invicta and a second, closely related, introduced species, S. richteri. The native fire ant, S. geminata, is genetically the most distinct of the three species studied, in accord with its taxonomic placement in a different species complex. Hypotheses concerning the derivation of the polygyne form of S. invicta from the monogyne form which invoke their long-term reproductive isolation in South America and separate introductions to North America appear unfounded. Although S. invicta and S. richteri are known to hybridize in North America, our study provided no evidence of gene introgression between S. invicta and the native species, S. geminata, in areas where our samples were collected. Analyses of population structure in S. invicta failed to reveal significant differentiation of populations or local inbreeding. Levels of genetic diversity in the three species studied, although not significantly different, were in the order predicted from knowledge of the population biology and recent history of the taxa, with S. richteri exhibiting the least diversity, S. geminata the greatest, and S. invicta having an intermediate level.     

Fire ants: What do rural and urban areas show us about occurrence, diversity,and ancestral state reconstruction?

In South America, Solenopsis saevissima and S. invicta are the most common fire ants. Nests are founded in areas under anthropic interference like urban or rural areas, but S. invicta is found preferentially in those with the greatest anthropic interference. However, we do not know the rates at which they exist in anthropized areas next to high density of native vegetation. Areas with 60 to 90% of native Atlantic Forest were selected to verify the occurrence of both species in rural and urban areas. We investigated the molecular diversity and applied the reconstruction of the ancestral state analysis for each species. A total of 186 nests were analyzed and we found that the two species had the same proportion in the urban area. However, S. saevissima had a higher rate of prevalence in the rural area, in addition to having a greater number of haplotypes and ancestry associated with this type of habitat for the region. S. invicta had the same number of haplotypes in both rural and urban regions, and less haplotypic diversity. We conclude that S. saevissima is a species typically associated with rural areas and S. invicta, although present, is not dominant in urban areas. Keywords: Biodiversity, haplotype diversity, DNA barcoding, mtDNA     

Host preference in Pseudacteon phorid flies: response of P. tricuspis and P. curvatus to black, red and hybrid imported Solenopsis fire ants in multiple choice bioassays

Host preferences in both sexes of Pseudacteon tricuspis Borgmeier (Jaguariuna biotype) and Pseudacteon curvatus Borgmeier (Formosa biotype) and their relative attraction to the imported fire ants (IFA), Solenopsis invicta Buren (red IFA), Solenopsis richteri Forel (black IFA) and S. invicta × S. richteri hybrids (hybrid IFA) were investigated in two separate experiments utilizing multiple choice flight bioassays. The results of both experiments clearly showed that both sexes of the Jaguariuna biotype of P. tricuspis could distinguish among the three IFA species and demonstrated greater preference for hybrid IFA and red IFA. This conclusion is supported by a variety of data collected on the number of fly visits, attack rate, and hovering duration (Experiment 1), and on the number of trapped flies (Experiment 2), which showed that black IFA is the least preferred of the three species. Similar results were recorded for the Formosan biotype of P. curvatus, although the data were not as strongly conclusive. Females of this biotype spent a significantly greater amount of time in hovering mode over red IFA and hybrid IFA compared to black IFA, but the other data were not significant. The red IFA is the natural host of both phorid fly biotypes and our results suggest that both biotypes may have evolved a specialized relationship with red IFA including an ability to discriminate it from related fire ants. These results are discussed in relation to the possible role of fire ant chemicals in mediating host preferences in phorid flies, contributions of male phorid flies to fire ant biocontrol, and the practical implications of the key findings.     

Morphology and number of Ommatidia in the compound eyes of Solenopsis invicta, Solenopsis richteri, and their hybrid (Hymenoptera: Formicidae)

The morphology and number of ommatidia of the compound eyes of Solenopsis invicta, Solenopsis richteri and their hybrid are investigated. Comparisons are made of the morphology and number of ommatidia in S. invicta, S. richteri and their hybrid. Major workers have significantly more facets than minor workers, male reproductive more than female reproductive, and the reproductives more than workers. Workers and reproductives have significantly more ommatidia in S. invicta than in S. richteri and significantly fewer in the hybrid than in either species. The major workers of each species have significantly wider facets than the minor workers, and workers have significantly wider facets than reproductives, which showed no difference in their facet diameter. There are no significant differences when the facet diameter is compared between the members of the castes of different species. These morphological and numerical dissimilarities are discussed in connection with known behavioral differences in the castes.     

Evolutionary history of <Emphasis Type="Italic">Wolbachia</Emphasis> infections in the fire ant <Emphasis Type="Italic">Solenopsis invicta</Emphasis>

Background  

Wolbachia are endosymbiotic bacteria that commonly infect numerous arthropods. Despite their broad taxonomic distribution, the transmission patterns of these bacteria within and among host species are not well understood. We sequenced a portion of the wsp gene from the Wolbachia genome infecting 138 individuals from eleven geographically distributed native populations of the fire ant Solenopsis invicta. We then compared these wsp sequence data to patterns of mitochondrial DNA (mtDNA) variation of both infected and uninfected host individuals to infer the transmission patterns of Wolbachia in S. invicta.     

A case study of human exacerbation of the invasive species problem: transport and establishment of polygyne fire ants in Tallahassee, Florida, USA

Understanding how exotic invasive species are spread is fundamental for ecology and conservation biology. Human transport has become one of the primary modes of dispersal for exotic species. We examined how the multiple queen, or polygyne, social form of the fire ant Solenopsis invicta is spread along roadsides in Tallahassee, Florida, USA. We then determined the likely source of this expanding population, which was a central soil depot. A survey of road maintenance practices in counties of several southeastern states and Texas revealed that the use of a central soil depot is a common practice. Road maintenance therefore may be the primary source for the establishment of new polygyne fire ant populations in this region and elsewhere. Control efforts focused on the soil depots will help to limit further spread of polygyne fire ants and perhaps other invasive organisms, particularly invasive weeds.     

A South American fire ant,Solenopsis nr. saevissima,in Guadeloupe,French West Indies

Several species of fire ants, well known for their painful stings, have spread from their native ranges and have become invasive pests, notably Solenopsis geminata, Solenopsis invicta, and Solenopsis richteri. Here, I report the first known exotic spread of another fire ant, Solenopsis nr. saevissima, a South American species now established on the French West Indian island of Guadeloupe. In 2008, I first found this shiny yellow fire ant at Plage de Viard beach on Guadeloupe. Genetic analyses indicated that this species was close to the yellow–brown variant of S. saevissima known only from Western Amazonia, but not an exact match. I returned to Guadeloupe in 2011 and found S. nr. saevissima at Plage de Viard plus five additional sites along a >20 km stretch on the east coast of Basse-Terre, from the Jarry commercial shipping port in the north to Plage Roseau in the south. It seems most likely that S. nr. saevissima arrived in Guadeloupe from South America in a commercial shipment unloaded at the port in Jarry. It is unclear whether this species will develop into a major new problem. Solenopsis nr. saevissima may simply replace another pernicious fire ant, S. geminata, which apparently arrived in Guadeloupe from South America several 100 years ago.     

Invasion history of Solenopsis invicta （Hymenoptera： Formicidae） in Fujian,China based on mitochondrial DNA and its implications in development of a control strategy

One of the most invasive species worldwide, Solenopsis invicta Buren, has been described in China since 2003. Recent studies have suggested that China populations are the result of introductions from the USA; however, detailed molecular studies need to be performed in order to understand the expansion and potential multiple introductions from other countries into China. As there were populations of red imported fire ant, S. invicta in different areas and with different methods of introduction, mitochondrial cytochrome oxidase Ⅰ gene was used as a marker from 12 populations in four cities in Fujian Province, China, to determine the relationship of invasion among these populations. The three most common haplotypes previously describe in invasive populations of S. invicta： H5, H22 and H36, were found in Fujian. However, frequencies in each city were different. For instance, three populations from Longyan city which invaded with waste plastics, shared haplotype H5. Populations from Xiamen city and Jinjiang city which dispersed with nursery stock, sward and scrap leather, shared haplotype H22. The population from Nanyan village of Xinluo district, Longyan city, bore haplotype H36. Mitochondrial data reveals that the invasion history of S. invicta in Fujian Province is complex, including multiple invasions probably from other provinces within China. Security measures to prevent S. invicta spreading within China are as important as from overseas.     

BIOCHEMICAL PHENOTYPIC AND GENETIC STUDIES OF TWO INTRODUCED FIRE ANTS AND THEIR HYBRID (HYMENOPTERA: FORMICIDAE)

Two introduced fire ants, Solenopsis invicta and S. richteri, and their hybrid were studied using phenotypic markers from gas chromatographic analysis of hydrocarbons and venom alkaloids, as well as genetic markers from enzyme electrophoresis. Both methods show that extensive gene introgression is occurring over a distance of at least 120 km at the contact zone between the two forms in eastern Mississippi. Genetic analyses suggest that the hybrid population does not depart dramatically from panmixia. Also, recombinant genotypes predominate in the hybrid zone, indicating that F₁ hybrids are viable. Allele frequency clines through the hybrid zone are apparent for four polymorphic loci. Data sets generated by the chromatographic and electrophoretic methods are highly concordant in that they differentiate completely between the two forms and agree in designating colonies from the contact zone as hybrid or parental in a high proportion (90%) of cases. The two methods can serve as complementary tools for studying closely related but genetically distinct populations in this, and perhaps other, groups of insects.     

Post-Release Host-Specificity Testing of Pseudacteon tricuspis, a Phorid Parasitoid of Solenopsis Invicta Fire Ants

Inherent in any biological control program is the risk of nontarget effects. Pseudacteon tricuspisBorgmeier, a parasitoid phorid fly, has been introduced to the United States from South America as a potential biocontrol agent of the red imported fire ant, Solenopsis invictaBuren. We conducted tests of host specificity on introduced populations of P. tricuspis, which are attracted to alarm pheromones released by their hosts during events such as mound disturbances and interspecific interactions. We monitored disturbed mounds of S. invicta and its close congener, S. geminata(F.), during the expansion of P. tricuspis across north Florida and after populations had been established for ~3 years. We also tested host acceptance in established populations of P. tricuspis by offering trays containing S. invicta, S. geminata, and 14 additional ant species representing 12 different non-Solenopsis genera. Although P. tricuspiswas commonly observed to hover over and attempt to oviposit on S. invicta, we never observed any parasitization attempts on any other ant species. As predicted by laboratory tests, released populations of P. tricuspis appear to be highly host specific and pose no obvious threat to nontarget species.     

Ecological dominance of the red imported fire ant, <Emphasis Type="Italic">Solenopsis invicta</Emphasis>, in its native range

Despite the widespread impacts invasive species can have in introduced populations, little is known about competitive mechanisms and dominance hierarchies between invaders and similar taxa in their native range. This study examines interactions between the red imported fire ant, Solenopsis invicta, and other above-ground foraging ants in two habitats in northeastern Argentina. A combination of pitfall traps and baits was used to characterize the ant communities, their dominance relationships, and to evaluate the effect of phorid flies on the interactions. Twenty-eight ant species coexisted with S. invicta in a gallery forest gap, whereas only ten coexisted with S. invicta in a xerophytic forest grassland. S. invicta was the most numerically dominant species in the richest and complex habitat (gallery forest); however it performed better as discoverer and dominator in the simpler habitat. S. invicta was active during day and night. In spite of its poor capacity to discover resources, S. invicta showed the highest ecological dominance and the second-best behavioral dominance after Camponotus blandus. S. invicta won 78% of the interactions with other ants, mostly against its most frequent competitor, Pheidole cf. obscurithorax, dominating baits via mass recruitment and chemical aggression. P. cf. obscurithorax was the best food discoverer. S. invicta won 80% of the scarce interactions with Linepithema humile. Crematogaster quadriformis was one of the fastest foragers and the only ant that won an equal number of contests against S. invicta. The low presence of phorid flies affected the foraging rate of S. invicta, but not the outcome of interspecific interactions. This study revealed that the red imported fire ant ecologically dominated other terrestrial ants in its native range; however, other species were able to be numerically dominant or co-dominant in its presence.     

Fire ant alarm pheromone and venom alkaloids act in concert to attract parasitic phorid flies, Pseudacteon spp.

Pseudacteon tricuspis, Pseudacteon obtusus and Pseudacteon curvatus are three species of parasitic phorid flies (Diptera: Phoridae), which have been introduced as classical biological control agents of imported, Solenopsis fire ants (Hymenoptera: Formicidae) in the southern USA. Previous studies demonstrated the behavioral response of P. tricuspis to the venom alkaloids and alarm pheromone of the fire ant, S. invicta. In the present study, we compared the responses of P. tricuspis, P. obtusus and P. curvatus to Solenopsis invicta alarm pheromone, venom alkaloids, or a mixture of both chemicals in four-choice olfactometer bioassays. The main hypothesis tested was that the fire ant alarm pheromone and venom alkaloids act in concert to attract Pseudacteon phorid flies. Both sexes of all three Pseudacteon species were attracted to low doses of the fire ant alarm pheromone or venom alkaloids (i.e. 1 ant worker equivalent) alone. However, the flies were significantly more attracted to a mixture of both chemicals (i.e., 1:1 mixture of alarm pheromone + alkaloids) than to either chemical. The results suggest an additive rather than a synergistic effect of combining both chemicals. Comparing the fly species, P. tricuspis showed relatively greater attraction to cis alkaloids, whereas the alkaloid mixture (cis + trans) was preferred by P. obtusus and P. curvatus. In general, no key sexual differences were recorded, although females of P. tricuspis and P. obtusus showed slightly higher response than conspecific males to lower doses of the alarm pheromone. The ecological significance of these findings is discussed, and a host location model is proposed for parasitic phorid flies involving the use of fire ant alarm pheromone and venom alkaloids as long range and short range attractants, respectively.     

Diet-induced nonmelanized cuticle in workers of the imported fire ant Solenopsis invicta buren

Nonmelanized cuticle development was induced in workers of Solenopsis invicta by feeding them an insect-free diet. The nonmelanized workers weighed less and had smaller mean headwidths than workers from normal colonies. Although nonmelanized ant colonies appeared to function normally in the laboratory, their attempts at stinging were felt only as “pin pricks.” Chemical analysis of venom alkaloids and cuticular hydrocarbons indicated no qualitative differences between nonmelanized and normal workers. Tyrosine, an essential amino acid tanning precursor, was found in adequate quantities in the free amino acid pool of nonmelanized ants. The specific cause of the nonmelanized condition is not known.     

Phytochemical differentiation of Galanthus nivalis and Galanthus elwesii (Amaryllidaceae): A case study

The alkaloid patterns of two occasionally sympatric Galanthus nivalis and Galanthus elwesii populations were studied by GC/MS. Thirty-seven alkaloids were detected, 25 for G. nivalis and 17 for G. elwesii. Only five alkaloids were found to occur in both species. The populations of Galanthus differed in their alkaloid biosynthetic pathways. Thus, the alkaloid pattern of G. nivalis was dominated by compounds coming from a para–para′ oxidative coupling of O-methylnorbelladine. The predominant alkaloids in the roots of this species were found to belong to the lycorine and tazettine structural types; bulbs were dominated by tazettine, leaves by lycorine and flowers by haemanthamine type alkaloids. In contrast, the alkaloid pattern of G. elwesii was dominated mainly by compounds coming from an ortho–para′ oxidative coupling. The predominant alkaloids in G. elwesii roots, bulbs and leaves were those of homolycorine type, whereas the flowers accumulated mainly tyramine type compounds. The chemotaxonomical value of the alkaloids found in the studied species is discussed.     

Molecular diversity of the microsporidium Kneallhazia solenopsae reveals an expanded host range among fire ants in North America

Kneallhazia solenopsae is a pathogenic microsporidium that infects the fire ants Solenopsis invicta and Solenopsis richteri in South America and the USA. In this study, we analyzed the prevalence and molecular diversity of K. solenopsae in fire ants from North and South America. We report the first empirical evidence of K. solenopsae infections in the tropical fire ant, Solenopsis geminata, and S. geminata × Solenopsis xyloni hybrids, revealing an expanded host range for this microsporidium. We also analyzed the molecular diversity at the 16S ribosomal RNA gene in K. solenopsae from the ant hosts S.invicta, S. richteri, S. geminata and S. geminata × S. xyloni hybrids from North America, Argentina and Brazil. We found 22 16S haplotypes. One of these haplotypes (WD_1) appears to be widely distributed, and is found in S. invicta from the USA and S. geminata from southern Mexico. Phylogenetic analyses of 16S sequences revealed that K. solenopsae haplotypes fall into one of two major clades that are differentiated by 2-3%. In some cases, multiple K. solenopsae haplotypes per colony were found, suggesting either an incomplete homogenization among gene copies within the 16S gene cluster or multiple K. solenopsae variants simultaneously infecting host colonies.     

Predicted range expansion of the invasive fire ant, Solenopsis invicta, in the eastern United States based on the VEMAP global warming scenario

The red imported fire ant, Solenopsis invicta Buren, is an invasive pest from South America that currently occupies much of the south‐eastern USA. Global warming is likely to allow range expansion of many invasive species, including S. invicta. We used a dynamic, ecophysiological model of fire ant colony growth coupled with models simulating climate change to predict the potential range expansion of S. invicta in the eastern USA over the next century. The climate change scenario predicted by the Vegetation–Ecosystem Modelling and Analysis Project (VEMAP) was used in our analyses. Our predictions indicate that the habitable area for S. invicta may increase by c. 5% over the next 40–50 years (a northward expansion of 33 ± 35 km). As the pace of global warming is expected to quicken in the latter half of the century, however, the habitable area for S. invicta in 2100 is predicted to be > 21% greater than it currently is (a northward expansion of 133 ± 68 km). Because the black imported fire ant, S. richteri Forel, occupies higher latitudes than S. invicta, the overall area of the eastern USA infested with invasive Solenopsis species could be greater than that estimated here.